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Principles of terrestrial ecosystem ecology.pdf

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50 3. Geology and Soils<br />

Phosphorus content<br />

Primary P<br />

races have very low productivity. The phenolic<br />

compounds produced by these trees as defenses<br />

against herbivores also retard decomposition,<br />

further reducing soil fertility (Northup et al.<br />

1995) (see Chapter 13).<br />

Potential Biota<br />

Total phosphorus<br />

Occluded<br />

Secondary P P<br />

Organic P<br />

Time<br />

Figure 3.4. The generalized effects <strong>of</strong> long-term<br />

weathering and soil development on the distribution<br />

and availability <strong>of</strong> phosphorus (P). Newly exposed<br />

geologic substrate is relatively rich in weatherable<br />

minerals, which release phosphorus. This release<br />

leads to accumulation <strong>of</strong> both organic and readily<br />

soluble forms (secondary phosphorus, such as<br />

calcium phosphate). As primary minerals disappear<br />

and secondary minerals capable <strong>of</strong> sorbing phosphorus<br />

accumulate, an increasing proportion <strong>of</strong> the<br />

phosphorus remaining in the system is held in<br />

unavailable (occluded) forms. Availability <strong>of</strong> phosphorus<br />

to plants peaks relatively early in this<br />

sequence and declines thereafter. (Redrawn with<br />

permission from Geoderma, Vol. 15 © 1976 Elsevier<br />

Science; Walker and Syers 1976.)<br />

The past and present organisms at a site<br />

strongly influence soil chemical and physical<br />

properties. Most soil development occurs in<br />

the presence <strong>of</strong> live organisms (Ugolini and<br />

Spaltenstein 1992). There are <strong>of</strong>ten clear associations<br />

between vegetation and soils. The<br />

organic acids in the litter <strong>of</strong> many coniferous<br />

species, for example, acidify the soil. This, in<br />

combination with the characteristically low<br />

quality <strong>of</strong> conifer litter, leads to slower decomposition<br />

in conifer than in deciduous forests<br />

(Van Cleve et al. 1991) (see Chapter 7). It is<br />

frequently difficult, however, to separate the<br />

chicken from the egg. Did the vegetation determine<br />

soil properties or vice versa?<br />

One approach to determining vegetation<br />

effects on soils has been to plant monocultures<br />

or species mixes into initially homogeneous<br />

sites. Rapidly growing grasses in a nitrogenpoor<br />

perennial grassland enhanced the nitrogen<br />

mineralization <strong>of</strong> soils within 3 years<br />

(Wedin and Tilman 1990) (see Fig. 12.5), as<br />

did deep-rooted forbs in an annual grassland<br />

(Hooper and Vitousek 1998). Another approach<br />

has been to examine the consequences<br />

<strong>of</strong> species invasions or extinctions on soil<br />

processes. The invasion <strong>of</strong> a non-native nitrogen<br />

fixer into Hawaiian rain forests, for<br />

example, increased nitrogen inputs to the<br />

system more than fivefold, altering the characteristics<br />

<strong>of</strong> soils and the colonization and competitive<br />

balance among native plant species<br />

(Vitousek et al. 1987) (see Fig. 12.3).<br />

Animals also influence soil properties. Earthworms,<br />

termites, and invertebrate shredders<br />

strongly influence decomposition rates (see<br />

Chapter 7) and therefore soil properties that<br />

are influenced by soil organic content.<br />

Human Activities<br />

Since the 1950s, the tripling <strong>of</strong> the human<br />

population and associated agricultural and<br />

industrial activities have strongly influenced<br />

soil development worldwide. Human activities<br />

influence soils directly through changes in<br />

nutrient inputs, irrigation, alteration <strong>of</strong> soil<br />

microenvironment through land use change<br />

(see Chapter 14), and increased erosional loss<br />

<strong>of</strong> soils. Human activities indirectly affect soils<br />

through changes in atmospheric composition<br />

and through the deletions and additions <strong>of</strong> species.<br />

Today and in the future, human activities<br />

will affect <strong>ecosystem</strong> properties both directly<br />

and through their effects on other interactive<br />

controls (see Chapters 14 and 16).<br />

Controls over Soil Loss<br />

Soil formation depends on the balance between<br />

deposition, erosion, and soil development. Soil<br />

thickness varies with hillslope position, with

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